Summary• Different portions of tree root systems play distinct functional roles, yet precisely how to distinguish roots of different functions within the branching fine-root system is unclear.• Here, anatomy and mycorrhizal colonization was examined by branch order in 23 Chinese temperate tree species of both angiosperms and gymnosperms forming ectomycorrhizal and arbuscular-mycorrhizal associations.• Different branch orders showed marked differences in anatomy. First-order roots exhibited primary development with an intact cortex, a high mycorrhizal colonization rate and a low stele proportion, thus serving absorptive functions. Second and third orders had both primary and secondary development. Fourth and higher orders showed mostly secondary development with no cortex or mycorrhizal colonization, and thus have limited role in absorption. Based on anatomical traits, it was estimated that c. 75% of the fine-root length was absorptive, and 68% was mycorrhizal, averaged across species.• These results showed that: order predicted differences in root anatomy in a relatively consistent manner across species; anatomical traits associated with absorption and mycorrhizal colonization occurred mainly in the first three orders; the single diameter class approach may have overestimated absorptive root length by 25% in temperate forests.

We have limited understanding of architecture and morphology of fine root systems in large woody trees. This study investigated architecture, morphology, and biomass of different fine root branch orders of two temperate tree species from Northeastern China-Larix gmelinii Rupr and Fraxinus mandshurica Rupr -by sampling up to five fine root branch orders three times during the 2003 growing season from two soil depths (i.e., 0-10 and.10-20 cm). Branching ratio (R b ) differed with the level of branching: R b values from the fifth to the second order of branching were approximately three in both species, but markedly higher for the first two orders of branching, reaching a value of 10.4 for L. gmelinii and 18.6 for F. mandshurica. Fine root diameter, length, SRL and root length density not only had systematic changes with root order, but also varied significantly with season and soil depth. Total biomass per order did not change systematically with branch order. Compared to the second, third and/or fourth order, the first order roots exhibited higher biomass throughout the growing season and soil depths, a pattern related to consistently higher R b values for the first two orders of branching than the other levels of branching. Moreover, the differences in architecture and morphology across order, season, and soil depth between the two species were consistent with the morphological disparity between gymnosperms and angiosperms reported previously. The results of this study suggest that root architecture and morphology, especially those of the first order roots, should be important for understanding the complexity and multi-functionality of tree fine roots with respect to root nutrient and water uptake, and fine root dynamics in forest ecosystems.

Root diameter, a critical indicator of root physiological function, varies greatly among tree species, but the underlying mechanism of this high variability is unclear. Here, we sampled 50 tree species across tropical and temperate zones in China, and measured root morphological and anatomical traits along the first five branch orders in each species. Our objectives were (i) to reveal the relationships between root diameter, cortical thickness and stele diameter among tree species in tropical and temperate forests, and (ii) to investigate the relationship of both root morphological and anatomical traits with divergence time during species radiation. The results showed that root diameter was strongly affected by cortical thickness but less by stele diameter in both tropical and temperate species. Changes in cortical thickness explained over 90% of variation in root diameter for the first order, and ∼74-87% for the second and third orders. Thicker roots displayed greater cortical thickness and more cortical cell layers than thinner roots. Phylogenetic analysis demonstrated that root diameter, cortical thickness and number of cortical cell layers significantly correlated with divergence time at the family level, showing similar variation trends in geological time. The results also suggested that trees tend to decrease their root cortical thickness rather than stele diameter during species radiation. The close linkage of variations in root morphology and anatomy to phylogeny as demonstrated by the data from the 50 tree species should provide some insights into the mechanism of root diameter variability among tree species.

Tree roots are highly heterogeneous in form and function. Previous studies revealed that fine root respiration was related to root morphology, tissue nitrogen (N) concentration and temperature, and varied with both soil depth and season. The underlying mechanisms governing the relationship between root respiration and root morphology, chemistry and anatomy along the root branch order have not been addressed. Here, we examined these relationships of the first- to fifth-order roots for near surface roots (0-10 cm) of 22-year-old larch (Larix gmelinii L.) and ash (Fraxinus mandshurica L.) plantations. Root respiration rate at 18 °C was measured by gas phase O2 electrodes across the first five branching order roots (the distal roots numbered as first order) at three times of the year. Root parameters of root diameter, specific root length (SRL), tissue N concentration, total non-structural carbohydrates (starch and soluble sugar) concentration (TNC), cortical thickness and stele diameter were also measured concurrently. With increasing root order, root diameter, TNC and the ratio of root TNC to tissue N concentration increased, while the SRL, tissue N concentration and cortical proportion decreased. Root respiration rate also monotonically decreased with increasing root order in both species. Cortical tissue (including exodermis, cortical parenchyma and endodermis) was present in the first three order roots, and cross sections of the cortex for the first-order root accounted for 68% (larch) and 86% (ash) of the total cross section of the root. Root respiration was closely related to root traits such as diameter, SRL, tissue N concentration, root TNC : tissue N ratio and stele-to-root diameter proportion among the first five orders, which explained up to 81-94% of variation in the rate of root respiration for larch and up to 83-93% for ash. These results suggest that the systematic variations of root respiration rate within tree fine root system are possibly due to the changes of tissue N concentration and anatomical structure along root branch orders in both tree species, which provide deeper understanding in the mechanism of how root traits affect root respiration in woody plants.

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